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![Titanodula fruhstorferi comb. nov., additional specimen ♂ [AS1 -Tam Dao N.P.] (RBINS). A, habitus, dorsal view. B, habitus, ventral view. C, head, frontal view. D, left prothoracic leg, anterior view. Titanodula fruhstorferi comb. nov., holotype ♀ (NHMW). E, habitus, dorsal view. F, habitus, ventral view. G, detail of head and prothoracic legs. H, labels. © A−D, Xavier Vermeersch and E−H, Harald Bruckner)](profile/Xavier-Vermeersch/publication/343280783/figure/fig3/AS:918540844556288@1596008611068/Titanodula-fruhstorferi-comb-nov-additional-specimen-AS1-Tam-Dao-NP-RBINS-A.png)
Titanodula fruhstorferi comb. nov., additional specimen ♂ [AS1 -Tam Dao N.P.] (RBINS). A, habitus, dorsal view. B, habitus, ventral view. C, head, frontal view. D, left prothoracic leg, anterior view. Titanodula fruhstorferi comb. nov., holotype ♀ (NHMW). E, habitus, dorsal view. F, habitus, ventral view. G, detail of head and prothoracic legs. H, labels. © A−D, Xavier Vermeersch and E−H, Harald Bruckner)
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![Fig. 2. Titanodula attenboroughi sp. nov., paratype ♀, [PT1] (RBINS)....](profile/Xavier-Vermeersch/publication/343280783/figure/fig1/AS:918540840357890@1596008610783/Titanodula-attenboroughi-sp-nov-paratype-PT1-RBINS-A-habitus-dorsal-view-B_Q320.jpg)
Recent taxonomic expeditions that were made possible within the framework of the Global Taxonomic Initiative project “A step further in the entomodiversity of Vietnam” resulted in the collecting of large and robust Hierodula-like praying mantises with a unique morphology of the male genitalia for which the new genus Titanodula gen. nov. is created....
Citations
... The Mantodea family Mantidae is currently composed of 76 genera worldwide (Schwarz & Roy, 2019;Vermeersch, 2020;Shcherbakov & Vermeersch, 2020). Many of the giant praying mantises from the oriental region with a typical praying mantis appearance are currently placed in the tribe Hierodulini of the subfamily Hierodulinae (Vermeersch, 2020). ...
... The Mantodea family Mantidae is currently composed of 76 genera worldwide (Schwarz & Roy, 2019;Vermeersch, 2020;Shcherbakov & Vermeersch, 2020). Many of the giant praying mantises from the oriental region with a typical praying mantis appearance are currently placed in the tribe Hierodulini of the subfamily Hierodulinae (Vermeersch, 2020). Hierodula is, however, up to this day a taxonomically very problematic genus, with a somewhat ambiguous 'one fits all' generic diagnosis. ...
... Although many Hierodula species look similar to each other superficially based on their external morphology, their male genitalia structures can be vastly different between species and geographic distribution. How all these species are phylogenetically related to each other remains somewhat unclear and requires further in-depth investigation (Vermeersch, 2020;Vermeersch & Unnahachote, 2020;Wang et al., 2020;Liu et al., 2021). Phylogenetic relationships of the whole genus and other related genera should be revised using modern, including molecular, tools to obtain a more natural taxonomy for this complex group of praying mantises. ...
Two new species of giant praying mantises, Hierodula kaltenbachi sp. nov. and Hierodula insperata sp. nov. from Northern and Southern Thailand respectively, are described based on recently collected material. A description of the habitus of both sexes, male genitalia, and photographs of live and mounted specimens are provided for both new species. The placement of the new species within the genus Hierodula is briefly discussed. A distribution map is also provided.
... An extremely diverse and wide-ranging genus in desperate need of thorough taxonomic revision (Schwarz & Konopik 2014;Vermeersch 2020). Seven species have been recorded from Australian territories, of which four are known to definitely occur: H. majuscula (Tindale, 1923) (Queensland), H. patellifera (Audinet-Serville, 1838) (Christmas Island), H. pustulifera (Wood-Mason, 1878) (Queensland), and H. werneri (Giglio-Tos, 1912) (Western Australia, Northern Territory, and Queensland). ...
Despite recent advances in Mantodean systematics that have resolved many historical problems within the order, the Australian fauna still contains many erroneously recorded or incorrectly placed species. An investigation of these problematic species and related taxa necessitated a number of taxonomic changes: Exparoxypilus Beier is a new synonym of Paraoxypilus Saussure, Paraoxypilus verrauxii Wood-Mason and Exparoxypilus africanus Beier are new synonyms of Paraoxypilus verreauxii Saussure, Pthersigena Hinton is a new synonym of Phthersigena Stål, Austromantis alboterminata Sjöstedt is a new synonym of Austromantis albomarginata Sjöstedt, Deiphobe australiana Giglio-Tos is a new synonym of Coenomantis kraussiana Saussure, Notomantis Tindale is a new synonym of Pseudomantis Saussure, Hierodula dimorpha Werner is a new synonym of Pseudomantis albofimbriata (Stål), Notomantis chlorophana Tindale is a new synonym of Pseudomantis hartmeyeri Werner, Pseudomantis papua (Werner) comb. nov. is transferred from Hierodula, Orthodera marginata (Saussure) is reinstated as a valid species, and Ciulfina liturgusa Giglio-Tos and Orthodera gunnii Le Guillou are considered nomina dubia. In addition, notes are provided on the nomenclature, systematics, distribution, and ecology of several other Australian taxa, and a synonymic checklist of Australian Mantodea is provided. Finally, the origins and diversification of the Australian fauna are discussed in light of these and other recent changes.
... Other than online databases, published literature was also used to identify the Mantodean specimens. These include Beier (1942), Bragg (2010Bragg ( , 2021, Chatterjee & Mukherjee (2013), Ehrmann & Borer (2015), Ehrmann & Roy (2009), Giglio-Tos (1915, 1917, 1927, Liu et al. (2020), Liu et al. (2021), Norman & Mustaffa (2019), Oliveira (1996-7), Rehn (1903), Roy (2009Roy ( , 2010, Shcherbakov (2012Shcherbakov ( , 2017, Schwarz et al. (2018aSchwarz et al. ( , 2018b, Schwarz & Shcherbakov (2017), Svenson (2014), Svenson & Vollmer (2014), Svenson et al. (2015), Vermeersch (2020), Werner (1933), Westwood (1889), and Wieland (2008). Many of the old publications can be obtained from The Biodiversity Heritage Library (BHL 2021) and HathiTrust Digital Library (HathiTrust 2021). ...
... Hierodula is one of the hardest genera in Mantodea to undergo identification (Schwarz & Konopik 2014, Bragg 2021, especially when there are only written descriptions without proper illustration or measurements recorded (Bragg 2021: 30 & 32) and the taxonomy within this genus is problematic (Vermeersch 2020, Vermeersch & Unnahachote 2020. In addition, the locality of a few type specimens of Hierodula are unknown (Bragg 2021: 31), making the species harder to identify. ...
... The sdpm (median secondary distal process) which is characteristic of Titanodula is absent, and the sdpl (lateral secondary distal process) of this specimen is long and straight, rather than the short and curved process that is found in Titanodula spp. (Vermeersch 2020, Liu et al. 2021. ...
Praying mantis curation and revision were conducted at the Forest Research Centre, Sepilok (FRCS), which led to the uncovering of at least 39 species of Mantodea. Identification was based on published literature and online databases, and the results were compiled into an annotated checklist for FRCS. There are five Bornean endemic species, one newly recorded species (Statilia apicalis), and one confirmed species for Borneo (Bimantis malaccana) in the FRCS collection. Photos were taken for each species in FRCS collection.
... At the same time, Tani et al. (2021) pointed out that the identification of Hierodula species recorded in GenBank needs to be reconsidered. In fact, the species H. formosana was classified into genus Titanodula in a taxonomic revision of giant Oriental praying mantises (Vermeersch 2020). The precise characteristics of the male genitalia, especially the shape of the secondary distal process, of H. chinensis in this study (Fig. 3a-c) differ greatly from those of the male genitalia of species in genus Titanodula in generally and of T. formosana in particular. ...
We identified an alien praying mantis found in Japan as the giant Asian mantis Hierodula chinensis by observing the characteristics of male genitalia. Nymphs hatched from oothecae collected in Nara Prefecture, Japan, were reared in a breeding room and compared with nymphs of the native Japanese species H. patellifera. Different from H. patellifera, the H. chinensis individuals passed through 5–7 nymphal instars before emerging as adults. The body color of the H. chinensis nymphs was brownish gray in the first instar, whereas the second and later instars were green. Unlike H. chinensis, H. patellifera had mottled patterns all over its body during the first to third instar. The first to third instar nymphs of H. chinensis had mottled patterns on the femurs of their forelegs, which is a diagnostic characteristic that no native praying mantis species has. From the fourth instar, 7–10 dorsal spines were observed on the foreleg coxa of H. chinensis, whereas only 3 or 4 spines were observed in H. patellifera. These findings show that H. chinensis and H. patellifera can be distinguished in all nymphal instars. We identified an alien praying mantis found in Japan as the giant Asian mantis Hierodula chinensis by observing the characteristics of male genitalia. Photographs of the genitalia of an adult male collected in Yoshino Town, Nara Prefecture, central Japan are shown. These major characteristics of the male genitalia matched those of H. chinensis from China and Korea, and those of H. sp. in Japan.
... Schwarz & Roy (2019) diagnosed Hierodulinae and listed two tribes (Archimantini and Hierodulini) assigning 23 genera to this subfamily. Recently, two new genera, Dracomantis Shcherbakov &Vermeersch, 2020, andTitanodula Vermeersch, 2020, were added to Hierodulinae (Shcherbakov & Vermeersch 2020;Vermeersch 2020), bringing the total in this subfamily to 25 genera with about 200 species (Otte et al. 2020). ...
... Schwarz & Roy (2019) diagnosed Hierodulinae and listed two tribes (Archimantini and Hierodulini) assigning 23 genera to this subfamily. Recently, two new genera, Dracomantis Shcherbakov &Vermeersch, 2020, andTitanodula Vermeersch, 2020, were added to Hierodulinae (Shcherbakov & Vermeersch 2020;Vermeersch 2020), bringing the total in this subfamily to 25 genera with about 200 species (Otte et al. 2020). ...
... For instance, two genera, Rhombomantis Ehrmann &Borer, 2015 andTitanodula Vermeersch, 2020, were established to include species previously included in Rhombodera or Hierodula. Current studies have shown that the male genitalia in Hierodulinae are highly diverse, strongly indicating the polyphyletic nature of several genera in this subfamily (Vermeersch 2020). Genital characters may be used to resolve phylogenetic relationships to a certain degree. ...
Species known from China in the praying mantis subfamily Hierodulinae are revised. A new species, Titanodula menglaensis sp. nov. is described. Hierodula tenuidentata Saussure, 1869 and Dracomantis mirofraternus Shcherbakov & Vermeersch, 2020 are newly recorded from China. Two new synonyms are proposed: Titanodula formosana (Giglio-Tos, 1912) = Titanodula fruhstorferi (Werner, 1916), syn. nov. and Hierodula macrodentata Wang, Zhou & Zhang, 2020, syn. nov. = Hierodula latipennis Brunner de Wattenwyl, 1893. Ootheca and male genitalia of the Chinese species are described and photographed. An identification key to genera and species of Hierodulinae from China is also provided. The current Chinese checklist contains 21 species.
... The oriental genus Hierodula Burmeister, 1838 is a very diverse and taxonomically problematic genus in need of complete revision (VERMEERSCH, 2020;SCHWARZ & ROY, 2019). The type species of the genus, Hierodula membranacea Burmeister, 1838, was described by BURMEISTER (1838) in a very brief fashion, mentioning two males stored at "Königl. ...
Hierodula confusa sp. nov. is described and figured based on material from Thailand. Habitus images of both sexes and photographs of the dissected male genitalia are given. Additionally, the male genitalia are compared with those of Hierodula membranacea Burmeister, 1838 and Rhombodera extensicollis Serville, 1839 and figured. The new species shows great similarity in external morphology with H. membranacea, which is the type species of the genus. However, it can be readily distinguished from H. membranacea by the morphology of the male genitalia, which have sufficiently different and constant characters across specimens within the known distribution range to justify the description of a new species. Given the similarity in their external morphology, H. confusa sp. nov. could easily have been misidentified as H. membranacea in the past, thus requiring a reassessment of collection material attributed to the species. R. extensicollis also shares morphological characters with the new species and is briefly discussed. A distribution map with confirmed records for Hierodula confusa sp. nov. is provided.
Species distribution models (SDMs) have conventionally been used for evaluating the distribution of individual species, but they can also be used, through comparing different SDMs, to evaluate the geographic similarity between taxa. In this study, we used a parasite and host system to infer the geographic overlaps between species with tight biological interaction, for example, parasites and their obligate host. Specifically, we used the horsehair worm Chordodes formosanus and its three mantis hosts to study the extent of niche overlap. We retrieved presence points for the host species and the parasite, and then we built SDMs with MaxEnt implemented in ENMeval using selected bioclim variables (based on variance inflation factor values) at 30s scale. The models showed that the hosts and parasite do not occur in the high elevation areas in Taiwan, which is expected based on their biology. Interestingly, the predicted parasite distribution included areas without collection records, implying local extinction or sampling bias. We subsequently evaluated niche overlap between hosts and the parasite according to five similarity indices (Schoener's D, I statistic, relative rank, Pearson correlation coefficient, and the rank correlation coefficient rho). Our models showed a high similarity of SDM predictions between hosts and the parasite. There were differences among metrics for which host shared the highest similarity with the parasite, but the majority of the results indicated that the Japanese boxing mantis had the highest niche similarity with the horsehair worm. The choice of the niche overlap metric to use can uncover information on the parasite's ecology, which can be important for endangered species. SDMs are reliable tools for host and parasite conservation management and could help improve our understanding of parasite biology and ecology.
This article aims to present a phylogenetic evaluation of Mantidae based on a mitochondrial genome (mitogenome) data set. The mitogenome of 10 Mantidae species were sequenced using next-generation sequencing. The length of nine the complete mitogenomes ranged from 15,371 bp in Tenodera aridifolia to 16,063 bp in Hierodula longa. Mantidae mitogenomes have 37 genes and control region with two exceptions: five trnR copies in Statilia maculata, and H. zhangi was incomplete missing trnI, trnQ, trnM and a portion of the control region. There was a large noncoding region (LNC) between trnM and nad2 in H. chinensis, H. longa, H. maculata and Titanodula sp. Most of protein-coding genes (PCGs) used the typical start ATN codon and TAA/TAG stop codons. All tRNAs fold into the typical clover-leaf secondary structure except trnS1 which lacks a dihydrouracil (DHU) arm. Nucleotide diversity and Ka/Ks analysis of 13 PCGs showed that atp8 had the highest variability and fastest evolutionary rate. Phylogenetic relationships among 42 Mantidae species were reconstructed using the 13 PCGs and two rRNA genes using Bayesian Inference (BI) and Maximum Likelihood (ML) methods. Of the seven mantid subfamilies included in this analysis, only four had multiple exemplars, and of those only Mantinae and Vatinae formed monophyletic groups in BI and ML trees. Consistent with previous studies, the monophyly of the Hierudulinae and Tenoderinae were not been supported. The present results imply that it is necessary to combine nuclear molecular markers and external characteristic to understand the phylogenetic relationships within Mantidae.
